Flagellated bacteria on nutrient-rich substrates can differentiate into a swarming state and move in dense swarms across surfaces. A recent experiment measured the flow in the fluid around an Escherichia coli swarm (Wu, Hosu and Berg, 2011 Proc. Natl. Acad. Sci. USA 108 4147). A systematic chiral flow was observed in the clockwise direction (when viewed from above) ahead of the swarm with flow speeds of about $10~μ$m/s, about 3 times greater than the radial velocity at the edge of the swarm. The working hypothesis is that this flow is due to the action of cells stalled at the edge of a colony that extend their flagellar filaments outwards, moving fluid over the virgin agar. In this work we quantitatively test his hypothesis. We first build an analytical model of the flow induced by a single flagellum in a thin film and then use the model, and its extension to multiple flagella, to compare with experimental measurements. The results we obtain are in agreement with the flagellar hypothesis. The model provides further quantitative insight into the flagella orientations and their spatial distributions as well as the tangential speed profile. In particular, the model suggests that flagella are on average pointing radially out of the swarm and are not wrapped tangentially.
Bacteriophage viruses, one of the most abundant entities in our planet, lack the ability to move independently. Instead, they crowd fluid environments in anticipation of a random encounter with a bacterium. Once they land on the cell body of their victim, they are able to eject their genetic material inside the host cell. Many phage species, however, first attach to the flagellar filaments of bacteria. Being immotile, these so-called flagellotropic phages still manage to reach the cell body for infection, and the process by which they move up the flagellar filament has intrigued the scientific…
Bacterial flagellar motor (BFM) is one of the ion-driven molecular machines, which drives the rotation of flagellar filaments and enable bacteria to swim in viscous solutions. Understanding its mechanism is one challenge in biophysics. Based on previous models and inspired by the idea used in description of motor proteins, in this study one two-state model is provided. Meanwhile, according to corresponding experimental data, mathematical relationship between BFM membrane voltage and pH value of the environment, and relationship between internal and external sodium concentrations are given. The…
In a classic paper, Edward Purcell analysed the dynamics of flagellated bacterial swimmers and derived a geometrical relationship which optimizes the propulsion efficiency. Experimental measurements for wild-type bacterial species E. coli have revealed that they closely satisfy this geometric optimality. However, the dependence of the flagellar motor speed on the load and more generally the role of the torque-speed characteristics of the flagellar motor is not considered in Purcell's original analysis. Here we derive a tuned condition representing a match between the flagella geometry and the …
Many species of bacteria swim through viscous environments by rotating multiple helical flagella. The filaments gather behind the cell body and form a close helical bundle, which propels the cell forward during a "run". The filaments inside the bundle cannot be continuously actuated, nor can they easily unbundle, if they are tangled around one another. The fact that bacteria can passively form coherent bundles, i.e. bundles which do not contain tangled pairs of filaments, may appear surprising given that flagella are actuated by uncoordinated motors. In this article, we establish the theoretic…
The growth of bacterial flagellar filaments is a self-assembly process where flagellin molecules are transported through the narrow core of the flagellum and are added at the distal end. To model this situation, we generalize a growth process based on the TASEP model by allowing particles to move both forward and backward on the lattice. The bias in the forward and backward jump rates determines the lattice tip speed, which we analyze and also compare to simulations. For positive bias, the system is in a non-equilibrium steady state and exhibits boundary-induced phase transitions. The tip spee…
The presence of a nearby boundary is likely to be important in the life cycle and evolution of motile flagellate bacteria. This has led many authors to employ numerical simulations to model near-surface bacterial motion and compute hydrodynamic boundary effects. A common choice has been the method of images for regularized Stokeslets (MIRS); however, the method requires discretization sizes and regularization parameters that are not specified by any theory. To determine appropriate regularization parameters for given discretization choices in MIRS, we conducted dynamically similar macroscopic …
In this supporting information we briefly describe the torque-speed measurement procedure. We show the hook spring compliance used in the simulations. We analyze the distribution functions of the moving and waiting time intervals. We study the dependence of the torque plateau region on the stator jumping rate and the cutoff angle, and the robustness of the results against different rotor-stator- potential and load-rotor forces.
The bacterial flagellar motor is an ion-powered transmembrane protein complex which drives swimming in many bacterial species. The motor consists of a cytoplasmic 'rotor' ring and a number of 'stator' units, which are bound to the cell wall of the bacterium. Recently, it has been shown that the number of functional torque-generating stator units in the motor depends on the external load, and suggested that mechanosensing in the flagellar motor is driven via a 'catch bond' mechanism in the motor's stator units. We present a method that allows us to measure -- on a single motor -- stator unit dy…
The bacterial flagellar motor is a highly efficient rotary machine used by many bacteria to propel themselves. It has recently been shown that at low speeds its rotation proceeds in steps [Sowa et al. (2005) Nature 437, 916--919]. Here we propose a simple physical model that accounts for this stepping behavior as a random walk in a tilted corrugated potential that combines torque and contact forces. We argue that the absolute angular position of the rotor is crucial for understanding step properties, and show this hypothesis to be consistent with the available data, in particular the observati…
Recent experiments on the bacterial flagellar motor have shown that the structure of this nanomachine, which drives locomotion in a wide range of bacterial species, is more dynamic than previously believed. Specifically, the number of active torque-generating units (stators) was shown to vary across applied loads. This finding invalidates the experimental evidence reporting that limiting (zero-torque) speed is independent of the number of active stators. Here, we propose that, contrary to previous assumptions, the maximum speed of the motor is not universal, but rather increases as additional …
Peritrichous bacteria synchronize and bundle their flagella to actively swim while disruption of the bundle leads to tumbling. It is still not known whether the number of flagella represents an evolutionary adaptation towards optimizing bacterial navigation. Here, we study the swimming dynamics of differentially flagellated Bacillus subtilis strains in a quasi-two-dimensional system. We find that decreasing the number of flagella N reduces the average turning angle between two successive run phases and enhances the duration and directional persistence of the run phase. As a result, having less…
The bacterial flagellar motor (BFM) is responsible for driving bacterial locomotion and chemotaxis, fundamental processes in pathogenesis and biofilm formation. In the BFM, torque is generated at the interface between transmembrane proteins (stators) and a rotor. It is well-established that the passage of ions down a transmembrane gradient through the stator complex provides the energy needed for torque generation. However, the physics involved in this energy conversion remain poorly understood. Here we propose a mechanically specific model for torque generation in the BFM. In particular, we i…
The bacterial flagellar motor plays a crucial role in both bacterial locomotion and chemotaxis. Recent experiments reveal that the switching dynamics of the motor depends on the motor rotation speed, and thus the motor torque, non-monotonically. Here we present a unified mathematical model which models motor torque generation based on experimental torque-speed curves and torque-dependent switching based on the conformational spread model. The model successfully reproduces the observed switching rate as a function of the rotation speed, and provides a generic physical explanation independent of…
University of Pennsylvania（宾夕法尼亚大学）；National Institute for Mathematical Sciences（韩国数学科学国立研究所）；Chung-Ang University（中央大学）；University of Cincinnati（辛辛那提大学）
A tiny bacterial flagellar motor rotates in both counter-clockwise (CCW) and clockwise (CW) rotational directions. The most important measurable characteristic of the flagellar motor is its torque versus angular speed relationship in CCW or CW modes, which is found to be non-symmetrical with each other, and still, such a phenomenon is not clearly understood.Here, we explain this asymmetry through a mechanistic model based on the detailed torque analysis for the rotation of the motor and the revolutionary as well as spinning motion of the filament and bead. We find out that the asymmetry result…
Trajectories and conformations of uni- and multiflagellar bacteria are studied with a coarse-grained model of a cell comprised of elastic flagella connected to a cell body. The elasticities of both the hook protein (connecting cell body and flagellum) and flagella are varied. Flexibility plays contrasting roles for uni- and multiflagellar swimmers. For a uniflagellar swimmer, hook and/or flagellar buckling occurs above a critical flexibility relative to the torque exerted by the flagellar motor. Addition of a second flagellum greatly expands the parameter regime of stable locomotion, because f…
Bacterial flagellar motor is one of the most complex and sophisticated nano machineries in nature. A duty ratio $D$ is a fraction of time that the stator and the rotor interact and is a fundamental property to characterize the motor but remains to be determined. It is known that the stator units of the motor bind to and dissociate from the motor dynamically to control the motor torque depending on the load on the motor. At low load where the kinetics such as a proton translocation speed limits the rotation rate, the dependency of the rotation rate on the number of stator units $N$ infers $D$; …
Most motile bacteria swim in viscous fluids by rotating multiple helical flagellar filaments. These semi-rigid filaments repeatedly join ('bundle') and separate ('unbundle'), resulting in a two-gait random walk-like motion of the cell. In this process, hydrodynamic interactions between the filaments are known to play an important role and can be categorised into two distinct types: direct interactions mediated through flows that are generated through the actuation of the filaments themselves, and indirect interactions mediated through the motion of the cell body (i.e. flows induced in the swim…
Switching of the direction of flagella rotations is the key control mechanism governing the chemotactic activity of E. coli and many other bacteria. Power-law distributions of switching times are most peculiar because their emergence cannot be deduced from simple thermodynamic arguments. Recently it was suggested that by adding finite-time correlations into Gaussian fluctuations regulating the energy height of barrier between the two rotation states, one can generate a power-law switching statistics. By using a simple model of a regulatory pathway, we demonstrate that the required amount of co…
The bacterial flagellar motor drives the rotation of flagellar filaments and enables many species of bacteria to swim. Torque is generated by interaction of stator units, anchored to the peptidoglycan cell wall, with the rotor. Recent experiments [Yuan, J. & Berg, H. C. (2008) PNAS 105, 1182-1185] show that near zero load the speed of the motor is independent of the number of stators. Here, we introduce a mathematical model of the motor dynamics that explains this behavior based on a general assumption that the stepping rate of a stator depends on the torque exerted by the stator on the rotor.…
To rotate continuously without jamming, the flagellar filaments of bacteria need to be locked in phase. While several models have been proposed for eukaryotic flagella, the synchronization of bacterial flagella is less well understood. Starting from a reduced model of flexible and hydrodynamically-coupled bacterial flagella, we rigorously coarse-grain the equations of motion using the method of multiple scales, and hence show that bacterial flagella generically synchronize to zero phase difference via an elastohydrodynamic mechanism. Remarkably, the far-field rate of synchronization is maximiz…
Rotation of a single bacterial flagellar motor is powered by multiple stators tethered to the cell wall. In a "power-stroke" model the observed independence of the speed at low load on the number of stators is explained by a torque-dependent stepping mechanism independent of the strength of the stator tethering spring. On the other hand, in models that depend solely on the stator spring to explain the observed behavior, exceedingly small stator spring constants are required. To study the dynamics of the motor driven by external forces (such as those exerted by an optical tweezer), back-steppin…
Spontaneous oscillations and geometric cutoff in confined bacterial swarms
Center of Materials Science and Optoelectronics Engineering, College of Materials Science and Opto-Electronic Technology, University of Chinese Academy of Sciences (UCAS)（中国科学院大学材料科学与光电技术学院物质科学与光电工程中心）；Department of Physics, Hong Kong Baptist University（香港浸会大学物理系）；Center for Interdisciplinary Studies, Westlake University（西湖大学交叉研究中心）
Bacteria are perhaps the simplest living systems capable of complex behaviour involving sensing and coherent, collective behaviour an example of which is the phenomena of swarming on agar surfaces. Two fundamental questions in bacterial swarming is how the information gathered by individual members of the swarm is shared across the swarm leading to coordinated swarm behaviour and what specific advantages does membership of the swarm provide its members in learning about their environment. In this article, we show a remarkable example of the collective advantage of a bacterial swarm which enabl…
Most bacteria swim through fluids by rotating helical flagella which can take one of twelve distinct polymorphic shapes. The most common helical waveform is the "normal" form, used during forward swimming runs. To shed light on the prevalence of the normal form in locomotion, we gather all available experimental measurements of the various polymorphic forms and compute their intrinsic hydrodynamic efficiencies. The normal helical form is found to be the most hydrodynamically efficient of the twelve polymorphic forms by a significant margin - a conclusion valid for both the peritrichous and pol…
In nature, bacterial collectives typically consist of multiple species, which are interacting both biochemically and physically. Nonetheless, past studies on the physical properties of swarming bacteria were focused on axenic (single species) populations. In bacterial swarming, intricate interactions between the individuals lead to clusters, rapid jets and vortices that depend on cell characteristics such as speed and length. In this work, we show the first results of rapidly swarming mixed-species populations of {\sl Bacillus subtilis} and {\sl Serratia marcescens}, two model swarm species th…
A wide spectrum of Peritrichous bacteria undergo considerable physiological changes when they are inoculated onto nutrition-rich surfaces and exhibit a rapid and collective migration denoted as swarming. Thereby, the length of such swarmer cells and their number of flagella increases substantially. In this article, we investigated the properties of individual E. coli-type swarmer cells confined between two parallel walls via mesoscale hydrodynamic simulations, combining molecular dynamics simulations of the swarmer cell with the multiparticle particle collision dynamics approach for the embedd…
To initiate tumbling of an E. coli, one of the helical flagella reverses its sense of rotation. It then transforms from its normal form first to the transient semicoiled state and subsequently to the curly-I state. The dynamics of polymorphism is effectively modeled by describing flagellar elasticity through an extended Kirchhoff free energy. However, the complete landscape of the free energy remains undetermined because the ground state energies of the polymorphic forms are not known. We investigate how variations in these ground state energies affect the dynamics of a reversely rotated flage…
Propagating interfaces are ubiquitous in nature, underlying instabilities and pattern formation in biology and material science. Physical principles governing interface growth are well understood in passive settings; however, our understanding of interfaces in active systems is still in its infancy. Here, we study the evolution of an active-passive interface using a model active matter system, bacterial swarms. We use ultra-violet light exposure to create compact domains of passive bacteria within Serratia marcescens swarms, thereby creating interfaces separating motile and immotile cells. Pos…
Recent experiments proposed to use confined bacteria in order to generate flows near surfaces. We develop a mathematical and a computational model of this fluid transport using a linear superposition of fundamental flow singularities. The rotation of a helical bacterial flagellum induces both a force and a torque on the surrounding fluid, both of which lead to a net flow along the surface. The combined flow is in general directed at an angle to the axis of the flagellar filament. The optimal pumping is thus achieved when bacteria are tilted with respect to the direction in which one wants to m…
Models based on surfactant driven instabilities have been employed to describe pattern formation by swarming bacteria. However, by definition, such models cannot account for the effect of bacterial sensing and decision making. Here we present a more complete model for bacterial pattern formation which accounts for these effects by coupling active bacterial motility to the passive fluid dynamics. We experimentally identify behaviours which cannot be captured by previous models based on passive population dispersal and show that a more accurate description is provided by our model. It is seen th…
Many bacteria are motile by means of one or more rotating rigid helical flagella, making them the only known organism to use rotation as a means of propulsion. The rotation is supplied by the bacterial flagellar motor, a particularly powerful rotary molecular machine. At the base of each flagellum is the hook, a soft helical polymer that acts as a universal joint, coupling rotation of the rigid membrane-spanning rotor to rotation of the rigid extra-cellular flagellum. In multi-flagellated bacterial species, where thrust is provided by a hydrodynamically coordinated bundle of flagella, the flex…
Unraveling bacterial strategies for spatial exploration is crucial for understanding the complexity in the organization of life. Bacterial motility determines the spatio-temporal structure of microbial communities, controls infection spreading and the microbiota organization in guts or in soils. Most theoretical approaches for modeling bacterial transport rely on their run-and-tumble motion. For Escherichia coli, the run time distribution was reported to follow a Poisson process with a single characteristic time related to the rotational switching of the flagellar motors. However, direct measu…
Inclined surfaces bias bacterial swarming expansion and edge organization
Global College, Shanghai Jiao Tong University（上海交通大学全球学院）；School of Physics, Georgia Institute of Technology（佐治亚理工学院物理学院）；Department of Mechanical Engineering, Johns Hopkins University（约翰霍普金斯大学机械工程系）；Intelligent Medicine Institute, Shanghai Medical College, Fudan University（复旦大学上海医学院智能医学研究所）
Bacteria commonly live in structured communities that affect human health and influence ecological systems. Heterogeneous populations, such as motile and non-motile populations, often coexist in bacteria communities. Motile subpopulations in microbial communities are believed to be important to dispersal, quest for food, and material transport. However, except in circumstances where motile cells drive colony expansion (e.g. bacterial swarming), the physiological functions of motile subpopulations in bacterial communities are largely unclear. Here we discovered that motile cells in routinely cu…
Hydrodynamics and confinement dominate bacterial mobility near solid or air-water boundaries, causing flagellated bacteria to move in circular trajectories. This phenomenon results from the counter-rotation between the bacterial body and flagella and lateral drags on them in opposite directions due to their proximity to the boundaries. Numerous experimental techniques have been developed to confine and maneuver motile bacteria. Here, we report observations on Escherichia coli and Enterobacter sp. when they are confined within a thin layer of water around dispersed micrometer-sized particles sp…
Bacterial motility, and in particular repulsion or attraction towards specific chemicals, has been a subject of investigation for over 100 years, resulting in detailed understanding of bacterial chemotaxis and the corresponding sensory network in many bacterial species. For Escherichia coli most of the current understanding comes from the experiments with low levels of chemotactically-active ligands. However, chemotactically-inactive chemical species at concentrations found in the human gastrointestinal tract produce significant changes in E. coli's osmotic pressure, and have been shown to lea…
We study the chemotaxis of a population of genetically identical swimming bacteria undergoing run and tumble dynamics driven by stochastic switching between clockwise and counterclockwise rotation of the flagellar rotary system. Understanding chemotaxis quantitatively requires that one links the switching rate of the rotary system in a gradient of chemoattractant/repellant to experimental measures of the efficiency of a population of bacteria in moving up/down the gradient. Here we achieve this by using a probabilistic model and show that the response of a population to the gradient is complex…
Bacterial processes ranging from gene expression to motility and biofilm formation are constantly challenged by internal and external noise. While the importance of stochastic fluctuations has been appreciated for chemotaxis, it is currently believed that deterministic long-range fluid dynamical effects govern cell-cell and cell-surface scattering - the elementary events that lead to swarming and collective swimming in active suspensions and to the formation of biofilms. Here, we report the first direct measurements of the bacterial flow field generated by individual swimming Escherichia coli …
Cooperation is vital for the survival of a swarm$^1$. Large scale cooperation allows murmuring starlings to outmaneuver preying falcons$^2$, shoaling sardines to outsmart sea lions$^3$, and homo sapiens to outlive their Pleistocene peers$^4$. On the micron-scale, bacterial colonies show excellent resilience thanks to the individuals' ability to cooperate even when densely packed, mitigating their internal flow pattern to mix nutrients, fence the immune system, and resist antibiotics$^{5-14}$. Production of an artificial swarm on the micro-scale faces a serious challenge $\frac{\;\;}{\;\;}$ whi…
